REDEFINITION OF THE ELEODES ESCHSCHOLTZSUBGENERA TRICHELEODES BLAISDELL AND

PSEUDELEODES BLAISDELL, WITH THE DESCRIPTIONOF A NEW SPECIES (COLEOPTERA: TENEBRIONIDAE)

M. ANDREW JOHNSTON

School of Life Sciences, Arizona State University PO Box 874501 Tempe, AZ 85287-4501; ajohnston@asu.edu

Abstract.— The Eleodes Eschscholtz, 1829 subgenera Tricheleodes Blaisdell, 1909 andPseudeleodes Blaisdell, 1909 are reviewed and redefined, based largely on femaleovipositors. Eleodes hirsuta LeConte, 1861 is designated as the type species ofTricheleodes, which is redefined to contain only that species. Eleodes barbatus Wickham,1918 is placed as incertae sedis within Eleodes. Pseudeleodes is expanded to includeTrichoderulus Blaisdell, 1923 syn. nov., as well as the following species: Eleodescaudiferus LeConte, 1858, E. leechi Tanner, 1961, E. longipilosus Horn, 1891, E. pilosusHorn, 1870, E. spoliatus Blaisdell, 1933, and E. tribulus Thomas, 2005, all comb. nov.;and Eleodes inornatus Johnston sp. nov. Eleodes granosus LeConte, 1866 is recognizedas a single valid species, which includes Eleodes inyoensis Tanner, 1961 syn. nov. andEleodes granosa pilifera Boddy, 1957 syn. nov. Eleodes caudiferus LeConte is similarlyidentified as a single valid species, which includes E. caudifera forma glabra Blaisdell,1909 syn. nov., E. caudifera forma scabra Blasidell, 1909 syn. nov., and E. caudiferaforma sublaevis Blaisdell, 1909 syn. nov. Eleodes pilosus Horn is found to be a singlespecies including E. obesus Doyen, 1985 as a syn. nov. A key is presented for the speciesbelonging to the subgenus Pseudeleodes. The new classification is reconciled with thecumulative preceding taxonomic perspective according to Blaisdell (1909) throughTriplehorn and coauthors (2015), by representing both taxonomies and aligning theirrespective concepts with Region Connection Calculus (RCC-5) articulations. The resulting,logically consistent alignment is inferred and visualized using the Euler/X multi-taxonomyalignment toolkit.

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Key words.— Classification, darkling beetles, morphology, synonymy, systematics.

INTRODUCTION

The desert stink beetles of the genus EleodesEschscholtz, 1829, belonging to the tribe AmphidoriniLeConte, 1862, are highly diverse and have a tangledtaxonomic history (Thomas 2005, Johnston et al.2015). The last comprehensive revision of the genuswas undertaken by Blaisdell (1909) who delineatedsubgenera primarily based on characters of the female

ovipositor. Revisionary work since that time has most-ly focused on subgeneric revisions (Triplehorn andThomas 2011, Aalbu et al. 2012, Johnston 2015, Tri-plehorn et al. 2015, Triplehorn and Thomas 2015).Female ovipositor characters have continued to be theprimary character system used to delineate subgen-era (e.g., Tri plehorn and Thomas 2011, Johnston 2015)and are consistent with molecular analyses (Smith et al. in prep). In order to accurately place a recently

A N N A L E S Z O O L O G I C I (Warszawa), 2016, 66(4): 665-679

PL ISSN 0003-4541 © Museum and Institute of Zoology PASdoi: 10.3161/00034541ANZ2016.66.4.018

discovered species, described herein, clarification ofthe subgeneric concepts of Tricheleodes Blaisdell,1909 and Pseudeleodes Blaisdell, 1909 was required.The re sults of that study, informed heavily by the fe -male ovi positor, are presented here.

The subgenus Tricheleodes was erected by Blais-dell (1909) to hold the two species Eleodes hirsutusLeConte, 1861 and Eleodes pilosus Horn, 1870, whichare extremely similar in habitus. The species Eleodesbarbatus Wickham, 1918 and E. spoliatus Blaisdell,1933 were added to the subgenus by the respectiveauthors upon their description. Tanner (1961) addedEleodes leechi and provided a key to the species thenbelonging to the subgenus. Triplehorn (1975) providednew distributional records for Eleodes barbatus. The description of Eleodes obesus Doyen, 1985 addeda sixth species to Tricheleodes which has remaineduntreated since.

The subgenus Pseudeleodes was erected by Blais-dell (1909) to hold the single species Eleodes gra-nosus LeConte, 1866 and was hypothesized as closelyrelated to Tricheleodes based on the very similarshape of the coxite and gonostyle of the female ovipos-itor, but differed by the vestiture and sculpturing of theintegument (Blaisdell 1909). The subspecies Eleodesgranosus piliferus Boddy, 1957 was described for a pilose form found in Oregon (Boddy 1957). Tanner(1961) described Eleodes inyoensis Tanner, 1961 andprovided a key to the two species of the subgenus asrecognized at the time. In a checklist of North Ameri-can Tenebrionidae, Eleodes caudiferus glabrusBlaisdell 1909, E. caudiferus scabrus Blaisdell 1909,and E. caudiferus sublaevis Blaisdell 1909, original-ly described as infrasubspecific forms, were elevated to subspecific standing (Papp 1961, see also Thomas2005).

Female genitalia have been very important for tene-brionid systematics (Tschinkel and Doyen 1980, Iwanand Kaminski 2016). Within Amphidorini, the termfemale genitalia is often used when discussing theshape of the ovipositor and has not historically in -volved internal structures of the genitalic tract (e.g.,Blaisdell 1909, Triplehorn and Thomas 2011). A com-prehensive examination of all internal and externalstructures has proven informative at the genus level inother groups (Iwan and Kaminski 2016) and may be anuntapped set of characters within Amphidorini. Thestudy presented here builds on the previous work inthis tribe and focuses on external features of theovipositor. The female terminalia of Amphidorini (Fig.1A, B) are modified from the generalized tenebrionidterminalia (Tschinkel and Doyen 1980) possessingshort paraprocts and a coxite comprised of a singlelobe. The ovipositor characters used to delimit subgen-era are in the shape and orientation of the coxite aswell as the shape and position of the gonostyle (Fig. 1).

Variation in the form of paraprocts and spiculum ven-tralae have not been reported before and were notfound to be diagnostic within the scope of this study.

MATERIALS AND METHODS

In total, 330 specimens representing all 11 speciestreated herein were examined using a Leica MZ16stereomicroscope. Females (n = 40) representing allspecies but Eleodes leechi were dissected by removingthe 4th and 5th visible sternites from the pinned speci-mens, dissecting out the terminalia in warm water,which were then (when necessary) gently cleared in10% KOH at room temperature for 30 minutes beforebeing point mounted. Habitus and ovipositor dissectionimages were taken using either the Visionary DigitalPassport II or Visionary Digital BK systems, equippedwith a Cannon 7D camera and 65 mm lens. Terminolo-gy for ovipositor anatomy follow Iwan and Kaminski(2016). Type specimens for all species were examinedin-person at the California Academy of Sciences orthrough the Museum of Comparative Zoology TypeData base (available on-line, http://insects.oeb.harvard.edu/mcz/) for the LeConte and Horn species.

The following collection codens are used for deposi -tories of new type material:ASUHIC – Arizona State University, Hasbrouck Insect

Collection; Tempe, AZ;CASC – California Academy of Sciences; San Fran-

cisco, CA;MAJC – M. Andrew Johnston Collection; Tempe, AZ;RLAC – Rolf L. Aalbu Collection; Sacramento, CA;

WBWC – William B. Warner Collection; Chandler, AZ.

SYSTEMATICS

Eleodes subgenus Tricheleodes Blaisdell 1909

Eleodes subgenus Tricheleodes Blaisdell 1909: 138.

Type species. Eleodes hirsuta LeConte, 1861: 352,by present designation.

Diagnosis. The combination of entirely hirsutebody, femora lacking spines in both sexes, tarsi withplantar groove uninterrupted and lined with darkspicules, and transverse mentum will separate thissubgenus from the rest of Amphidorini, except forPseu deleodes and some members of LitheleodesBlaisdell 1909 (see Triplehorn and Thomas 2015). Tri -cheleodes can be separated from the latter by the fe -male ovipositor possessing an evenly triangular coxitewith the gonostyle mammiliforn and inserted apicallyon coxite (Fig. 1D), versus the coxite being subrectan-gular with a flattened, fleshy gonostyle in Lithe leodesor the coxite broadly triangular with a mammiliform

666 M. A. JOHNSTON

gonostyle inserted laterally along distal margin, clear-ly separated from the produced inner lobe of the coxitein Pseudeleodes (Fig. 1B).

Redescription. Body oblong-ovate, black (Fig. 2B).Mentum transverse, middle lobe arcuate, barely pro-jecting beyond lateral lobes. Pronotum subquadrate,arcuate laterally, densely punctured, each puncturebearing a single long seta. Elytra asperate to muricate,densely punctured, pilose. Anterior femora simple,without spines in both sexes. Anterior tibia with long itudinal carina along basal third of exterior face.Protarsi with basal segment thickened anteriorly in

females, unmodified in males, basal three segmentsbearing dorsal setae nearly as long as the followingsegment, plantar surface of all segments lined withblack spicules. Female ovipositor (Fig. 1D) with coxiteweakly to moderately sclerotized, subtriangular, innerlobe not produced apically, gonostyle mammiliform,positioned apically on dorsal face.

Discussion. The subgenus Tricheleodes, as rede-fined here, only includes the type species Eleodeshisutus (Fig. 2B). The form of the female genitalia isquite different from the remainder of the species for-merly treated in this subgenus which are described

REDEFINITION OF TRICHELEODES AND PSEUDELEODES 667

Figure 1. Eleodes ovipositors. (A) Eleodes (Pseudeleodes) tribulus Thomas ventral view, right paraproct, coxite, and gonostyle highlighted. (B) Eleodes (Pseudeleodes) tribulus Thomas dorsal view, right paraproct, coxite, and gonostyle highlighted. (C) Eleodes (incertae sedis) barbatus

Wickham, right coxite and gonostyle highlighted. (D) Eleodes (Tricheleodes) hirsutus LeConte, right coxite and gonostyle highlighted.

below. Eleodes hirsutus was described from the GreatSalt Lake in Utah (LeConte 1861) and has since beenfound across central and northern Nevada west intothe Owens Valley and Sierra Nevada mountains of California.

Eleodes Incertae sedis

Eleodes barbatus Wickham, 1918: 256.

Eleodes barbatus (Fig. 2A) is among the smallestof all Eleodes, rivaled only by E. connatus Solier, 1848(Thomas 2015), and inhabits sandy habitats in the four-corners region of Utah, Arizona, and New Mexico (Tri -ple horn 1975, Johnston et al. 2015). The original de -scription (Wickham 1918: 257) states that this species“looks decidedly like an Amphidora” but opted forplacement in the genus Eleodes based on the femaleovipositor. The ovipositor (Fig. 1C) is strongly sclero-tized, with the inner lobe of the coxite strongly pro-duced and excavate posterolaterally. Similar forms of the strongly sclerotized and produced inner lobe ofthe coxite are also present in the genera Neoba-phion Blaisdell, 1925, Trogloderus Leconte, 1879 and

Lariversius Blaisdell, 1947. The oviposters of Tri che -le odes and Pseudeleodes (Fig. 1A, B, D) are complete-ly different and lack the modified inner lobe of the coxite. The subgenera Metablapylis Blaisdell, 1909and Bla pylis Horn, 1870 possess strongly sclerotizedfe male genitalia, but lack extremely produced innerlobes. It is possible that Eleodes barbatus represents a unique lineage of Amphidorini from any describedthus far, but it may be a highly derived member of oneof the groups discussed above. Until a detailed study ofthese generic and subgeneric boundaries can be com-pleted, it is hereby placed as incertae sedis withinEleodes.

Eleodes subgenus Pseudeleodes Blaisdell, 1909

Eleodes subgenus Pseudeleodes Blaisdell, 1909: 146.

Type species. Eleodes ganosa LeConte, 1866, bymonotypy.

Diagnosis. Species in the subgenus Tricheleodescan be recognized by the combination of: the entirebody clothed in black setae, varying from minute to long and conspicuous; presence of a quadrate

668 M. A. JOHNSTON

Figure 2. (A) Eleodes (incertae sedis) barbatus Wickham, syntype. (B) Eleodes (Tricheleodes) hirsutus LeConte. Scale bar = 1 cm.

men-tum; and elytra always with muricate to tubercu-late sculpturing and slightly divergent posterior apicallobes. Eleodes (Metablapylis) nigrinus LeConte,1858, Tricheleodes, and some Blapylis may be diffi-cult to separate from Pseudeleodes externally, andwill require examining female ovipositors (see below).

Redescription. Body oblong-ovate to elongate,black, always setose but varying widely from minuteand inconspicuous to erect and pilose. Mentum sub-quadrate, strongly arcuate anteriorly. Legs generallyslender and long. Anterior femora usually simple, occa-sionally with spines, sometimes sinuate apically. Ante-rior tibia with longitudinal carina along basal 1/2 to 3/4of exterior face. Protarsi with plantar surface linedwith dark spicules, never thickened or interrupted,basal three segments bearing dorsal setae extend-ing to distal margin of the following segment. Elytrafaintly muricate to strongly tuberculate, pseudepipleu-ron clearly extending to apex of elytra, strongly con-cave apically, caudate or not, elytral apicies slightlydivergent, appearing notched. Ovipositor (Fig. 3C, 4D)with coxite elongate, triangular at apex, inner lobestrongly produced, acute. Gonostyle positioned pos-terolaterally, not extending beyond apex of inner lobeof coxite.

Discussion. Since its original description by Blais-dell (1909: 147), Pseudeleodes has been presumed tobe closely related to Tricheleodes. The subgenericconcept for Pseudeleodes is herein expanded to ac -commodate the remainder of the species excluded fromTricheleodes as defined herein, several previouslyincertae sedis species (Triplehorn et al. 2015) as wellas an undescribed species. Members of Pseude leodesare frequently associated with sandy soils, and thishabitat specificity may drive the formation of disjunctpopulations. Investigation of external morphology andovipositors make several species-level classificatorychanges appropriate at this time. However, furtherphylogeographic and molecular phylogenetic studieson this subgenus may reveal cryptic species complexesand could provide insight into speciation patternsthroughout the Great Basin and related arid regions ofwestern North America.

Eleodes (Pseudeleodes) inornatus Johnston, sp. nov.(Fig. 3)

Diagnosis. This species can be distinguished by thefollowing combination of characters: body elongate,

REDEFINITION OF TRICHELEODES AND PSEUDELEODES 669

Figure 3. Eleodes (Pseudeleodes) inornatus Johnston, new species, scale bar = 1 cm. (A) Holotype female. (B) Paratype male. (C) Holotype female ovipositor, not to scale.

parallel sided, appearing glabrous but clothed inminute setae arising from punctures, anterior pronotalangles produced forward and acute, prosternumstrongly produced posteriorly of procoxae, acute apically, anterior femora sinuate in both sexes, elytrafinely muricate, punctures arranged in weak striae. It somewhat resembles Eleodes nigrinus LeConte,1858 in body shape, size, and sculpturing, but can bedistinguished by its prominent pronotal angles whichare not projected forward in E. nigrinus.

Description. Female. (Fig. 3A) Body length 15–20mm, width 5–7 mm (n = 5). Body elongate, parallelsided, clothed in minute, inconspicuous setae. Anten-nae clavate, not attaining posterior margin of prono-tum, segments 3–7 at least as long as wide, segments 8–11 wider than long, terminal 3 segments slightly flattened. Mentum tri-lobed; mesal lobe subquadrate,evenly arcuate anteriorly; bearing longitudinal keelmesally, evenly sloping into foveae posterolaterally,each bearing several black setae. Ligula transver-sely cordate, bearing stiff golden setae distally. Maxil-lae heavily microsculptured; cardo bearing fine, scattered setae, stipes bearing fine, scattered setaeand several thick black setae; palpiger strongly pro-duced; first palp segment twice as long as second, dis-tal segment subtriangular, apical width subequal tolength. Mandibles bifid apically; densely puncturedbearing stiff setae dorsally and basally. Labrum trans-verse, deeply notched anteriorly; weak transverseridge dorsally, densely setose anterad of ridge, finelysetose posterad. Head evenly punctured; punc-tures muricate, each bearing a minute seta. Gula evenly triangular, rufo piceus. Eyes vertical, weaklyemarginate.

Pronotum slightly wider than long; anterior anglesproduced, acute; lateral margin arcuate, slightly sinu-ate posteriorly; disc finely muricate and punctured,becoming granulate laterally, each puncture bearingminute seta. Propleura granulately tuberculate anteri-orly, each tubercle bearing one seta; smooth to slightlywrinkled posteriorly. Prosternum granulately tubercu-late anteriorly, becoming rugose between coxae; pro -sternal process extended well posteriorly of procoxae,strongly acute apically.

Scutellum transverse, evenly arcuate posteriorly;strongly microsculptured, lacking punctures. Elytrawith muricate punctures, fused medially; appearingglabrous, surface strongly microsculptured; punctureseach bearing a minute seta; punctures generally ar -ranged in striae, striae unimpressed; elytral apicesslightly divergent, appearing notched. Pseudepipleuronnot expanded anteriorly, narrowing slightly posterior-ly; becoming strongly concave along posterior half offifth visible abdominal sternite; pseudepipleural foldcontinuous until elytral apex. Meso- and meta-sternummoderately punctured.

Legs scabrously muricate. Fore femora clavate,weakly sinuate distally; more roughly sculptured ventrally than dorsally; slightly excavate anteriorly indistal 3/4. Fore tibia with longitudinal carina alongbasal half of exterior face; roughly tuburculate ventral-ly; tibial spurs similar, extending to distal margin ofsecond tarsal segment. Tarsi simple, clothed in darkblack setae; plantar surface uninterrupted, lined withblack spicules; basal 3 segments bearing dorsal setaeas long as proceeding segment; tarsal claws longerthan 5th tarsal segment. Middle and hind legs similar tofore legs except femora not sinuate apically.

Abdomen ventrally flat; 5 visible sternites, evenlyand sparsely covered with muricate punctures; inter-coxal process rectangular, broader than long; 5th visi-ble sternite more roughly punctured than precedingsternites; lined posteriorly with moderately long blacksetae.

Terminalia. (Fig. 2C) Coxite triangularly elongatedorsally, outer margin evenly tapering inward; innerlobe strongly produced distally; sparsely clothed inpale setae. Gonostyle mammiliform; inserted dorsolat-erally in depression distally on coxite; terminating wellbefore terminus of coxite, bearing tuft of setae, setaenearly as long as gonostyle.

Male. (Fig. 3B) Body length 14–16 mm, width 4–6mm (n = 3). Similar to female, slightly less robust, foretibia slightly more sinuate.

Terminalia. Adeagus cylindrical, slightly flattenedlaterally. Basal piece cylindrical, lateral margins sinu-ate basally, distal margin broadly and arcuately exca-vated. Parameres fused, elongately triangular dorsally;basal margin broadly arcuate; narrowly rounded dis-tally. Clavae with distal end of ventral surface rotatedoutwardly; slightly concave at distal tip, bearing goldensetae; apex broadly rounded.

Variation. Other than the range in body size notedabove, there is little variation in the specimens exam-ined for this new species. The setae can be very diffi-cult to observe in some portions of the elytra and onsome individuals. This may be due to individual varia-tion or perhaps to environmental wear to the cuticle inlonger-lived individuals.

Distribution. This species is only known fromregions of fine sandy soils from northern Nevada.

Etymology. Eleodes (Pseudeleodes) inornatus isso named for having the least noticeable sculpturingand vestiture of any known species of Pseudeleodes.

Remarks. This species has only been collected inthe month of July, and two of the eight known speci-mens were recently deceased upon collection. It is pos-sible that adults of this species are relatively shortlived. The specimens at the type locality in Chur-chill, Nevada, were found by flipping over bags of yardwaste dumped on the side of the road. The speci-mens from Teel’s Marsh dunes, Nevada, were found by

670 M. A. JOHNSTON

headlamping at night, including the two deceased spec-imens. In terestingly, Sand Mountain, Nevada, has beenfrequented by many coleopterists but only a single spec-imen has yet been identified from that locality, whichmay be considered as further evidence to the relativerarity and ephemerality of adult E. inornatus popula-tions.

Type material. Holotype (female) “USA: Nevada,Churchill/Co. 17mi N.Fallon 1190m/39.7098°,-118.733°/6.VII.2015 M.A. Johnston”, bearing a red label stating“HOLOTYPE/Eleodes inornatus Johnston 2016”deposited in CASC.

Paratypes. 7 specimens marked with blue paratypelabels stating “PARATYPE/Eleodes inornatus John-ston 2016”, 3 with same information as holotype 1 male,1 female (RLAC); 1 male (MAJC); “USA: Nevada, Min-eral/Co., Teel’s Marsh Dunes/38.211°, -118.307° 1500m/6.VII.2015 M.A. Johnston” 1 female, 1 male (ASUHIC);1 male (MAJC); “USA: NEV: Churchill Co./ Sand Mountain, 22mi./ SE of Fallon el. ~3940’/39°17’29”N,118°24’56”/July 1-8, 2011; barrier pit/falls, dune; W.B.Warner” 1 female (WBWC).

Checklist of Eleodes subgenus Pseudeleodes

Eleodes subgenus Pseudeleodes Blaisdell, 1909: 146.Eleodes subgenus Trichoderulus Blaisdell, 1923: 281, syn. nov.

Eleodes (Pseudeleodes) caudiferus LeConte,1858: 184, comb. nov.Eleodes caudifera forma glabra Blaisdell, 1909: 228, syn. nov.Eleodes caudifera forma scabra Blaisdell, 1909: 228, syn. nov.Eleodes caudifera forma sublaevis Blaisdell, 1909: 228, syn. nov.

Eleodes (Pseudeleodes) granosus LeConte, 1866:116.Eleodes inyoensis Tanner, 1961: 68, syn. nov.Eleodes granosa pilifera Boddy, 1957: 193, syn. nov.

Eleodes (Pseudeleodes) leechi Tanner, 1961: 63,comb. nov.

Eleodes (Pseudeleodes) longipilosus Horn, 1891:42, comb. nov.

Eleodes (Pseudeleodes) pilosus Horn, 1870: 314,comb. nov.Eleodes obesus Doyen, 1985: 232, syn. nov.

Eleodes (Pseudeleodes) inornatus Johnston, sp.nov.

Eleodes (Pseudeleodes) spoliatus Blaisdell, 1933:196, comb. nov.

Eleodes (Pseudeleodes) tribulus Thomas, 2005:552 [replacement name for Amphidora caudataHorn, 1870: 330], comb. nov.Trichoderulus longipilosus Blaisdell, 1923: 281, [syn. Triplehorn

and Aalbu 1987: 371].=Eleodes blaisdelli Doyen [in Doyen and Law rence], 1979: 367

[replacement name for T. longipilosis Blaisdell].

Both Blaisdell (1909:150) and Boddy (1957: 194)interpreted the species Eleodes granosus LeConte(Fig. 4) to be variable in sculpturing, with the elytraltubercles ranging from granulate to large and rounded.Thus Eleodes inyoensis Tanner (Fig. 4A) falls withinthe E. granosus concept of Blaisdell, and the holotypeof E. inyoensis shares the same sculpturing, thoughlacking erect setae, of the holotype of E. granosus pi-liferus Boddy (Fig. 4B). Analysis of types and speci-mens from across an extended geographic range,including specimens from Inyo County, California withlong setae (Fig. 4C), is most consistent with the recog-nition of a single species with variable ornamentationand fixed female ovipositor morphology (Fig. 4D). Thelength of elytral setae, pronotal punctation, and elytraltubercle sculpturing all seem to vary at the individuallevel and are not correlated with apparent geographicpatterns. Thus, both Eleodes inyoensis Tanner andE. granosus piliferus Boddy are hereby considerednew synonyms of E. granosus LeConte. This speciesis uncommon in collections and extends through manyecologically heterogeneous mountain ranges of easternCalifornia and southern Oregon and Idaho. It is hopedthat increased sampling of this striking species willlead to a more in-depth phylogeographic study in thefuture.

The enigmatic Eleodes tibulus Thomas, 2005(Figs. 5A,B) has a tangled taxonomic legacy, itselfbeing a replacement name for Amphidora caudataHorn, 1870 (Thomas 2005: 552). However, in accor-dance with present classification and synonymies, E. tribulus typifies the genus-group name Tricho -derulus Blaisdell, 1923 (Doyen and Lawrence 1979,Triplehorn and Aalbu 1987). This species has been dif-ficult to accommodate in the recognized subgenericdefinitions, and was therefore placed as incertae se -dis within Eleodes (Triplehorn and Aalbu 1987). Maleshave spines on the fore femora and caudate elytra,characters which are shared with some members of thenominate subgenus (e.g., Eleodes (Eleodes) spinipesSolier, 1848 and Eleodes (Eleodes) tenu ipes Casey,1890). The general facies of the species likens it toEleodes pilosus, herein placed in Pseu deleodes, aswell as the species formerly placed in Amphidorawhich were later moved to the subgenus Blapylis(Doyen and Lawrence 1979). Examination of the femaleovipositor (Figs. 1A,B, 5C), which possesses a triangu-lar coxite with a mammiliform gonostyle inserted

REDEFINITION OF TRICHELEODES AND PSEUDELEODES 671

apically on the outer lobe, indicates that E. tri bulusbelongs in Pseudeleodes Blaisdell. As such, the spe -cies is hereby transferred and results in Trichoderu-lus Blaisdell, 1923 being a new synonym of Pseude -leodes Blaisdell, 1909.

Eleodes caudiferus LeConte (Fig. 6A) and E. lon -gi pilosus Horn (Fig. 6B) have historically been treat-ed in the nominate Eleodes subgenus Eleodes (Blais-dell 1909, Tanner 1961) but were recently excludedupon revision (Triplehorn et al. 2015). Examination ofthe respective female ovipositors (Fig. 6C, D) indicatesthat the proper placement of these two species is inPseudeleodes. This placement of E. caudiferus is fur-ther supported by the study of Eleodes larvae (Smithet al. 2014). The two species are highly similar exter-nally, primarily distinguished by E. longipilosusbeing clothed in long setae, but have distinctly different

female genitalia (Fig. 6C,D) warranting specific status.In his monograph, Blaisdell (1909) described threeinfra-subspecific forms meant to diagnose inter-indi-vidual variation in elytral sculpturing. These formswere later elevated to subspecific status by Papp(1961). The taxonomic inconsistencies created by thislatter work have been discussed in Thomas (2005) andJohnston (2015). The adults of Eleodes caudiferusexhibit some variation in the rugosity of the elytralsculpturing, generally varying across a cline with spec-imens from the western extent of its range in Arizonatending to be more roughly sculptured than those fromeastern New Mexico and western Texas, the easternextent of the species range. Upon examination of spec-imens from across the species range, this seems to bemerely inter-individual variation, as originally pro-posed by Blaisdell (1909), and the three subspecific

672 M. A. JOHNSTON

Figure 4. Eleodes (Pseudeleodes) granosus LeConte diversity, scale bars = 1 cm. (A) Eleodes inyoensis Tanner holotype (syn of Eleodes(Pseudeleodes) granosus LeConte). (B) Eleodes granosus piluferus Boddy holotype (syn of Eleodes (Pseudeleodes) granosus LeConte).

(C) Hirsute form of Eleodes (Pseudeleodes) granosus. (D) Ovipositor, not to scale.

names are hereby placed in synonymy with Eleodescaudiferus.

Eleodes pilosus (Fig. 7A, C) is the most wide-spread species in the subgenus Pseudeleodes. Foundon sand dunes and in other sandy habitats from Cali-fornia’s Owens valley, north into Oregon, east toWyoming and south to western New Mexico, thisspecies would likely make an excellent system for a phylogeographic study. Other flightless tenebrionidshave shown high levels of genetic divergence acrosstheir range (Chatzimanolis and Caterino 2007, Smith etal. in prep.) and could represent cryptic species com-plexes depending on which species concept is applied.The evolutionary species concept of Wiley and Mayden

(2000) is applied here. Eleodes obesus as described byDoyen (1985) (Fig. 7C) is presumably distinguishedfrom E. pilosus by having shorter setae and femalesbeing more rotund. These characters alone are inade-quate to recognize separate species. Individuals of E. pilosus exhibit significant variation in vestiture,integument sculpturing, and body size yet they retainsimilarity in female ovipositors (Fig. 7B) throughouttheir range. The holotype female of E. obesus has a remarkably rotund abdomen, but more slenderfemales with the same shorter setal vestiture are foundthroughout the Sierra Nevada Mountains of California.Based on the lack of consistently co-varying charac-ters, E. obesus is hereby synonymized with E. pilosus.

REDEFINITION OF TRICHELEODES AND PSEUDELEODES 673

Figure 5. Scale bars = 1 cm. (A) Eleodes (Pseudeleodes) tribulus Thomas, male. (B) Eleodes (Pseudeleodes) tribulus Thomas, female. (C) Eleodes (Pseudeleodes) tribulus Thomas ovipositor, not to scale. D. Eleodes (Pseudeleodes) spoliatus Blaisdell, holotype.

Key to the species of Eleodes subgenusPseudeleodes Blaisdell

1. Elytra always caudate, fore femora never spined,body length > 15 mm . . . . . . . . . . . . . . . . . . . . . . . . 2

–. Elytra not caudate, or if so fore femora spined andbody < 15 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

2. Elytral setae short, inconspicuous (Fig. 6A); AZ,NM, UT . . . . . . . . . E. caudiferus LeConte (Fig. 6A)

–. Elytral setae long and pilose (Fig. 6B); northern NV . . . . . . . . . . . . . . E. longipilosus Horn (Fig. 6B)

3. Pronotal disc evenly tuberculate throughout . . . . . .. . . . . . . . . . . . . . . . E. spoliatus Blaisdell (Fig. 5D).

–. Pronotal disc minutely to rugosely punctate, occa-sionally tuberculate laterally . . . . . . . . . . . . . . . . . . 4

4. Elytra with fine, muricate punctures, setae incon-

spicuous . . . . . . . E. inornatus new species (Fig. 3)–. Elytra with long, conspicuous setae, or if not then

distinctly tuberculate . . . . . . . . . . . . . . . . . . . . . . . . 55. Pronotum minutely punctured, males caudate (Fig.

5A) . . . . . . . . . . . . . E. tribulus Thomas (Fig. 5A,B)–. Pronotum moderately to rugosely punctured, never

caudate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66. Elytra with large, rounded tubercles . . . . . . . . . . . .7–. Elytra finely muricate to granulately tuberculate . .

. . . . . . . . . . . . . . . . . . . . E. pilosus Horn (Fig. 7A,C)7. Pronotum very rugose, body length < 16 mm; four-

corners region of CO, UT . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . E. leechi Tanner (Fig. 7D)

–. Pronotum heavily punctate, body length > 15 mm;eastern Sierra Nevada region, CA, ID, NV, OR . . . .. . . . . . . . . . . . . . . . . . E. granosus LeConte (Fig. 4).

674 M. A. JOHNSTON

Figure 6. Scale bar = 1 cm. (A) Eleodes (Pseudeleodes) caudiferus LeConte, male. (B) Eleodes (Pseudeleodes) longipilosus Horn, female. (C) Eleodes (Pseudeleodes) caudiferus ovipositor, not to scale. (D) Eleodes (Pseudeleodes) longipilosus ovipositor, not to scale.

Classificatory changes represented as taxonomic concept alignment

Biological names continue to be integral to evolu-tionary and ecological research (Patterson et al. 2010),though integration of these names with other types ofbiological data is a major challenge (Page 2006, Patter-son et al. 2016). One problematic area is the use of thesame name string to reference incongruent taxonomicconcepts either through time (Franz et al. 2015), orrepresenting concurrent alternative taxonomic cir-cumscriptions (Franz et al. 2016a). A logical alignmentof two taxonomies provides both a useful visualiza-tion of taxonomic change and a logically consistent

framework for machine computation (Jansen andFranz 2015).

A consensus taxonomy representing the cumulativestatus of the herein treated taxonomic entities immedi-ately prior to this publication (Fig. 8A, “2015” taxono-my; representing Blaisdell 1909 through Triplehorn etal. 2015) was aligned to the newly proposed taxonomypresented here (Fig. 8A, “2016” taxonomy). A logicallyconsistent alignment was inferred using the Euler/Xtoolkit (Chen et al. 2014) following the methods ofFranz et al. (2016b), which takes as input both tax-onomies and a set of articulations (Fig. 8A, dashedlines) represented using region connection calculuswith five possible relationships (RCC-5; disconnected,

REDEFINITION OF TRICHELEODES AND PSEUDELEODES 675

Figure 7. Scale bars = 1 cm. (A) Eleodes (Pseudeleodes) pilosus LeConte, male. (B) E. (Pseudeleodes) pilosus female ovipositor, not to scale. (C) Eleodes obesus Doyen, holotype (syn of Eleodes (Pseudeleodes) pilosus LeConte). (D) Eleodes (Pseudeleodes) leechi Tanner, holotype.

676 M. A. JOHNSTON

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equal, partially overlapping, proper inclusion [A isincluded in B], and inverse proper inclusion [A in -cludes B]). The aligned taxonomies are visualized inFig. 8B. This visualization is particularly useful tointerpret congruent taxonomic concepts which mayspan multiple classificatory ranks, such as the 2015Pseudeleodes concept which had all of its memberssynonymized under the 2016 Eleodes granosus con-cept, thus making the two concepts equivalent. Thelarge number of incongruent 2015 taxonomic concepts(Fig. 8B, yellow octogons) is indicative of the moregranular classification, which was considered unten-able in 2016. The reassessment of types created manynomenclatural and classificatory changes in the 2016taxonomy (see the Pseudeleodes checklist above) pri-marily due to transferring species into Pseudeleodes.The taxonomic concept alignment, however, reinforcesthe consistency of species-level concepts throughoutthe changes between the two taxonomies. Raw inputand output files for the Euler/X alignment are availableon-line (https://github.com/mandrewj/PseudeleodesAlignment).

ACKNOWLEDGMENTS

Specimens were generously made available forstudy through Warren Steiner of the United StatesNational Museum, Matthew Gimmel of the Santa Bar-bara Natural History Museum, as well as from the per-sonal collections of William B. Warner, Ronald Somer-by, and Rolf Aalbu. Norman Penny and Anne Barberfrom the California Academy of Sciences made examin-ing specimens and imaging type material possible. NicoFranz assisted with the alignment and visualization oftaxonomies and along with Aaron Smith and PatriceBouchard provided valuable feedback on early versionsof this manuscript. This research was supported by the National Sciences Foundation (DEB–1258154/1523605), the Arizona State University EvolutionaryBiology Summer Research Fellowship, and the UnitedStates Department of Agriculture – Agricultural Re -search Service (Agreement 58-1275-1-335).

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Received: May 31, 2016Accepted: December 2, 2016